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Triassic Roots of the Amphiastraeid Scleractinian Corals

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Triassic Roots of the Amphiastraeid Scleractinian Corals J. Paleont., 75(1), 2001, pp. 34±45 Copyright q 2001, The Paleontological Society 0022-3360/01/0075-34$03.00 TRIASSIC ROOTS OF THE AMPHIASTRAEID SCLERACTINIAN CORALS EWA RONIEWICZ AND JAROSèAW STOLARSKI Instytut Paleobiologii, Polska Akademia Nauk, Twarda 51/55, 00-818 Warszawa, Poland, ,[email protected]., ,[email protected]. ABSTRACTÐThe Early Carnian (Upper Triassic) phaceloid coral originally described by Volz (1896) as Hexastraea fritschi, type species of Quenstedtiphyllia Melnikova, 1975, reproduced asexually by ``Taschenknospung'' (pocket-budding), a process documented herein for the ®rst time. This type of budding is recognized only in the Amphiastraeidae, a family thus far recorded only from Jurassic- Cretaceous strata. Similar to amphiastraeids, Quenstedtiphyllia fritschi (Volz, 1896) has separate septal calci®cation centers and a mid- septal zone built of serially arranged trabeculae. The most important discriminating characters of the new amphiastraeid subfamily Quenstedtiphylliinae are one-zonalendotheca and radial symmetry of the corallite in the adult stage (in contrast to two-zonal and bilateral symmetry in the adult stage in Amphiastraeinae). Quenstedtiphyllia fritschi shares several primitive skeletal characters (plesiomorphies) with representatives of Triassic Zardinophyllidae and, possibly, Paleozoic plerophylline rugosans: e.g., thick epithecal wall and strongly bilateral early blastogenetic stages with the earliest corallite having one axial initial septum. To interpret the phylogenetic status of amphiastraeid corals, we performed two analyses using plerophylline rugosans and the solitary scleractinian Protoheterastraea, respec- tively, as the outgroups. The resulting phylogenetic hypotheses support grouping the Zardinophyllidae with the Amphiastraeidae in the clade Pachythecaliina (synapomorphy: presence of pachytheca). Taschenknospung is considered an autapomorphy for the Amphiastraei- dae. This study is the ®rst attempt to analyze the relationships of the Triassic corals cladistically. INTRODUCTION Subsequent stages of the earliest phase of budding were traced at intervals of 2 mm due to a lack of material. CLERACTINIAN CORALS arose during the Middle/Late Triassic S as four major microstructural groups (Roniewicz and Mo- Institutional abbreviations. MGUWr, Geological Museum, rycowa, 1993; see also Fig. 1): 1) mini-trabecular (e.g., reimani- Wrocøaw University, Wrocøaw, Poland; ZPAL, Institute of Paleo- phylliids); 2) thick-trabecular (e.g., actinastreids, pamiroseriids); biology, Warsaw, Poland; GMH, Geiseltal Museum, Halle, Ger- 3) fascicular (5non-trabecular, i.e., stylophyllids and gigantostyl- many; UMIP, University of Montana, Institute of Paleontology, ids); and 4) pachythecal (zardinophyllids). Phylogenetic relation- USA. ships among these groups are poorly understood, but differences MORPHOLOGY OF THE SKELETON in microstructural characters, especially among pachythecal, sty- Zardinophyllids are solitary, i.e., Pachythecalis Cuif, 1975; lophyllid, and mini/thick-trabecular groups, suggests a polyphy- Zardinophyllum Montanaro-Gallitelli, 1975; and phaceloid, i.e., letic origin (Roniewicz and Morycowa, 1993). A polyphyletic or- Pachydendron Cuif, 1975; Pachysolenia Cuif, 1975 (probably the igin of Scleractinia is also suggested by phylogenetic analyses of senior subjective synonym of Lubowastraea Melnikova, 1986, a DNA sequences of shallow-water scleractinians (Romano and form with corallites apposed to each other). A very thick wall is Palumbi, 1996; Veron et al., 1996). a predominant skeletal character. Septa are non-exsert and usually Particularly intriguing is the origin and evolution of Triassic located deep in the calice. Adult coralla may have quasi-radial corals that have skeletal architecture strikingly similar to that of symmetry, whereas initial and juvenile stages are often strongly the late Paleozoic rugosans. One such coral group is the Zardi- bilaterally symmetrical. In solitary Pachythecalis and Zardino- nophyllidae, which resembles plerophylline rugosans by having phyllum, intracalicular space is ®lled during the ontogeny with an identical initial corallite ontogeny, an extremely thick corallite compact sclerenchyme (stereome), whereas in phaceloid Pachy- wall (in relation to the calice diameter), rhopaloid septa, and many solenia and Pachydendron this space is ®lled with tabuloid dis- other aspects of corallum morphology and microstructure (see sepiments. Cuif, 1975b; Cuif and Stolarski, 1999; Stolarski 1999). Zardino- Protoheterastraeids are restricted herein to solitary Protohet- phyllids are represented by four fossil genera: Pachythecalis Cuif, erastraea Wells, 1937, represented by the lectotype of the type 1975 (early Norian), Pachysolenia Cuif, 1975 (early Carnian± species Protoheterastraea leonhardi (Volz, 1896). Protoheteras- Norian), Pachydendron Cuif, 1975 (early Norian±Rhaetian), and traea has a relatively thick epithecal wall, tabuloid dissepiments, Zardinophyllum Montanaro-Gallitelli, 1975 (early Carnian± and radial symmetry. Cerioheterastrea, another genus assigned Rhaetian). Analysis of the skeletal structure and budding of the by Cuif (1976) to the ``Protoheterastraea-group'', differs signif- amphiastraeid Quenstedtiphyllia fritschi (Volz, 1896) has yielded icantly from Protoheterastraea in septal structure and endotheca, new data, which have encouraged us to renew discussion of the and in our opinion, these two genera should not be grouped to- origin and evolution of Zardinophyllidae. gether. Ladinian solitary/phaceloid corals with thick epitheca and tabular endotheca that resemble Protoheterastraea have been il- MATERIAL AND METHODS lustrated by Deng and Kong (1984, pl. 3, ®gs. 7, 9; identi®ed as Elasmophyllia and, respectively, Pinacophyllum). We examined the holotype of Quenstedtiphyllia fritschi (Volz, Amphiastraeids are solitary (Cheilosmilia Koby, 1888), pha- 1896), the only known specimen of this species. The specimen celoid (Amphiaulastrea Geyer, 1955; Aulastraea Ogilvie, 1897; (colony fragment, size 80 3 70 3 40 mm) was collected in the Mitrodendron Quenstedt, 1881; Pleurophyllia de Fromentel, lower Carnian St. Cassian Beds in the Dolomites (Southern Alps), 1856; Hexapetalum EliasÏova, 1975; Pseudopisthophyllum Geyer, and is housed at the Geiseltal Museum in Halle (Germany). Its 1955; Hykeliphyllum EliasÏova, 1975); cerioid (Pleurostylina de original aragonite mineralogy is preserved, thus enabling micro- Fromentel, 1856); or pseudocerioid with separated polygonal cor- structural and ontogenetic (blastogenetic) observations. We base allites (Amphiastraea Etallon, 1859). The corallites are usually our observations on thin sections (12 slides), acetate peels taken strongly bilaterally symmetrical. The wall is thick. The endotheca from transverse and longitudinal sections of the colony, and a is built of tabuloid dissepiments, which peripherally may pass into series of peels made every 0.1 mm from the budding corallite. a zone of vesicular dissepiments (Melnikova and Roniewicz, 34 RONIEWICZ AND STOLARSKIÐTRIASSIC AMPHIASTRAEID CORALS 35 FIGURE 1ÐFour basic skeletal microstructures of the Triassic Scleractinia. 1, Minitrabecular (e.g., Protoheterastraea). 2, thick-trabecular (e.g., Chondrocoenia) 3, fascicular (e.g., Stylophyllum). 4, pachythecal (e.g., Pachydendron). After Roniewicz (1996). 1976). For the present purposes, we have not included in this fascicles of ®bers with various dimensions to well delimited mod- analysis a group of Jurassic scleractinians traditionally classi®ed ules without axes (Fig. 2.1, 2.3). Generally, fascicles that form with amphiastraeids in the suborder Amphiastraeina Alloiteau, wall modules extend into the septa. 1952: i.e., Donacosmiliidae, Carolastraeidae, Intersmiliidae. They Septa.In zardinophyllids, calci®cation centers of the mid-sep- are known only from completely recrystallized specimens, and tal zone are, as a rule, not separated. They form a homogeneous are generally very poorly characterized, such that many of their zone composed of microcrystalline material in longitudinal and characters would be represented in a phylogenetic analysis as transverse sections of septa (vertical and horizontal sections are missing values, producing unreliable results. required to detect possibly horizontally, or respectively, vertically Quenstedtiphyllia fritschi (Volz) has a phaceloid growth form. arranged septal trabeculae). In transverse section, ®bers thicken- The corallites are radially symmetrical with penta- and hexameral ing the septum are often oriented strongly oblique to the mid- septal con®gurations composed of regularly distributed septa in 4 septal zone, indicating predominant axial accretion of the septum orders. The most striking morphological character is a thick wall (e.g., Pachythecalis major, Pachydendron microthallos, see Cuif, (Fig. 2). The calicular lumen is ®lled by tabuloid (one-zonal) en- 1975b, ®gs. 4, 6b respectively). Septal accretion was often rhyth- dotheca. mic, and its successive phases can be distinguished in transverse section as superimposed secretional units (e.g., in Pachythecalis MICROSTRUCTURE OF THE SKELETON Cuif, 1975b, ®g. 3b, d). To date, longitudinal sections of Pachy- Wall.Zardinophyllidae and Amphiastraeidae have a unique solenia cylindrica (type species of the genus) have not furnished type of epithecal wallÐa pachythecaÐwith an internal layer (i.e., us with microstructural details of septa. Thus, it is unclear whether epithecal, or rather pachythecal stereome) built of radially
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